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The Major Transitions in Evolution. Eörs Szathmáry. Collegium Budapest AND Eötvös University. Units of evolution. multiplication heredity variability. Some hereditary traits affect survival and/or fertility. The importance of cumulative selection.
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The Major Transitions in Evolution Eörs Szathmáry Collegium Budapest AND Eötvös University
Units of evolution • multiplication • heredity • variability Some hereditary traits affect survival and/or fertility
The importance of cumulative selection • Natural selection is a non-random process. • Evolution by natural selection is a cumulative process. • Cumulative selection can produce novel useful complex structures in relatively short periods of time.
John Maynard Smith (1920-2004) • Educated in Eaton • The influence of J.B.S. Haldane • Aeroplane engineer • Sequence space • Evolution of sex • Game theory • Animal signalling • Balsan, Kyoto, Crafoord prizes
The major transitions (1995) * * * * * These transitions are regarded to be ‘difficult’
The importance of cumulative selection • Natural selection is a non-random process. • Evolution by natural selection is a cumulative process. • Cumulative selection can produce novel useful complex structures in relatively short periods of time.
Difficulty of a transition • Selection limited (special environment) • Pre-emption: first come selective overkill • Variation-limited: improbable series of rare variations (genetic code, eukaryotic nucleocytoplasm, etc.)
Difficult transitions are ‘unique’ • Operational definition: all organisms sharing the trait go back to a common ancestor after the transition • These unique transitions are usually irreversible (no cell without a genetic code, no bacterium derived from a eukaryote can be found today)
Fisher’s(1930) question: the birth of ALife "No practical biologist interested in sexual reproduction would be led to work out the detailed consequences experienced by organisms having three or more sexes; yet what else should he do if he wishes to understand why the sexes are, in fact, always two?"
Units of evolution • multiplication • heredity • variation hereditary traits affecting survival and/or reproduction
Recurrent themesin transitions • Independently reproducing units come together and form higher-level units • Division of labour/combination of function • Origin of novel inheritance systems Increase in complexity • Contingent irreversibility • Central control
Hamilton’s rule b r> c • b:help given to recipient • r:degree of genetic relatedness between altruist and recipient • c:price to altruist in terms of fitness • Formula valid for INVASION and MAINTENANCE • APPLIES TO THE FRATERNAL TRANSITIONS!!!
Division of labour • Is advantageous, if the “extent of the market” is sufficiently large • If it holds that a “jack-of-all-trades” is a master of none • Not always guaranteed (hermaphroditism) • Morphs differ epigenetically
Most forms of multicellularity result from fraternal transitions • Cells divide and stick together • Economy of scale (predation, etc.) • Division of labour follows • Cancer is no miracle (Szent-Györgyi) • A main difficulty: “appropriate down-regulation of cell division at the right place and the right time” (E.S. & L. Wolpert)
What makes us human? • Note the different time-scales involved • Cultural transmission: language transmits itself as well as other things • A novel inheritance system
Fluid Construction Grammar with replicating constructs (with Luc Steels) • selective amplification by linked replication • mutation, recombination, etc.
Why is often no way back? • There are secondary solitary insects • Parthenogens arise again and again • BUT no secondary ribo-organism that would have lost the genetic code • No mitochondrial cancer • No parthenogenic gymnosperms • No parthenogenic mammals
Contingent irreversibility • In gymnosperms, plastids come from one gamete and mitochondria from the other: complementary uniparental inheritance of organelles • In mammals, so-called genomic imprinting poses special difficulties • Two simultaneous transitions are difficult squared: parthenogenesis per se combined with the abolishment of imprinting or complementary uniparental inheritance
Central control • Endosymbiotic organelles (plastids and mitochondria) lost most of their genes • Quite a number of genes have been transferred to the nucleus • The nucleus controls organelle division • It frequently controls uniparental inheritance, thereby reducing intragenomic conflict